Projections from the posterior intralaminar thalamic nuclei and the superior colliculus (SC) to the subthalamic nucleus (STN) and the zona incerta (ZI) have been described in the primate and rodent. The aims of this study was to investigate several questions on these projections, using modern neurotracing techniques in rats, to advance our understanding of the role of STN and ZI. We examined whether projection patterns to the subthlamus can be used to identify homologues of the primate centromedian (CM) and the parafascicular nucleus (Pf) in the rodent, the topography of the projection including what percent of intralaminar neurons participate in the projections, and electron microscopic examination of intralaminar synaptic boutons in STN. The aim on the SC-subthalamic projection was to examine whether STN is the main target of the projection. This study revealed: (i) the areas similar to primate CM and Pf could be recognized in the rat; (ii) the Pf-like area sends a very heavy topographically organized projection to STN but very sparse projection to ZI, which suggested that Pf might control basal ganglia function through STN; (iii) the projection from the CM-like area to the subthalamus was very sparse; (iv) Pf boutons and randomly sampled asymmetrical synapses had similar distributions on the dendrites of STN neurons; and (v) the lateral part of the deep layers of SC sends a very heavy projection to ZI and moderate to sparse projection to limited parts of STN, suggesting that SC is involved in a limited control of basal ganglia function.

Parvalbumin (PV)-positive interneurons form dendritic gap junctions with one another, but the connectivity among gap junction-coupled dendrites remains uninvestigated in most neocortical areas. We visualized gap junctions in layer 4 of the mouse barrel cortex and examined their structural details. PV neurons were divided into 4 types based on the location of soma and dendrites within or outside barrels. Type 1 neurons that had soma and all dendrites inside a barrel, considered most specific to single vibrissa-derived signals, unexpectedly formed gap junctions only with other types but never with each other. Type 2 neurons inside a barrel elongated dendrites outward, forming gap junctions within a column that contained the home barrel. Type 3 neurons located outside barrels established connections with all types including Type 4 neurons that were confined inside the inter-barrel septa. The majority (33/38, 86.8%) of dendritic gap junctions were within 75 μm from at least 1 of 2 paired somata. All types received vesicular glutamate transporter 2-positive axon terminals preferentially on somata and proximal dendrites, indicating the involvement of all types in thalamocortical feedforward regulation in which proximal gap junctions may also participate. These structural organizations provide a new morphological basis for regulatory mechanisms in barrel cortex.

The striatum is critically involved in execution of appropriate behaviors, but its internal structures remain unmapped due to its unique structural organization, leading to ambiguity when interpreting heterogeneous properties of striatal neurons that differ by location. We focused on site-specific diversity of striosomes/matrix compartmentalization to draw the striatum map. Five types of striosomes were discriminated according to diverse immunoreactivities for the µ-opioid receptor, substance P (SP) and enkephalin, and each type occupied a particular domain inside the striatum. Furthermore, there was an additional domain lacking striosomes. This striosome-free space was located at the dorsolateral region and received afferents preferentially from the primary motor and sensory cortices, whereas the striosome-rich part received afferents from associational/limbic cortices, with topography inside both innervations. The proportion of dopamine D1 receptor-expressing, presumptive striatonigral neurons was approximately 70% in SP-positive striosomes, 40% in SP-deficient striosomes, 30% in the striosome-free space, and 50% in the matrix. In contrast, the proportion of D2 receptor-expressing, presumptive striatopallidal neurons was complementary to that of D1 receptor-expressing cells, indicating a close relationship between the map and the direct and indirect parallel circuitry. Finally, the most caudal part of the striatum lacked compartmentalization and consisted of three lamina characterized by intense and mutually exclusive immunoreactivities for SP and enkephalin. This tri-laminar part also received specific afferents from the cortex. The newly obtained map will facilitate broad fields of research in the basal ganglia with higher resolution of the three-dimensional anatomy of the striatum.